Aposematic species have evolved conspicuous warning signals, such as bright colors and striking patterns, to deter predators. Some edible and harmless species take advantage of this deterrent effect by mimicking their appearance. Mimicry is a great example of how natural selection produces remarkable adaptations. However, while some species evolve a very close similarity to their models to effectively avoid attacks, others are successful in doing so despite an incomplete similarity, i.e. imperfect mimicry. In some cases, it is surprising how such a crude disguise can fool predators. Why and how imperfect mimicry can persist has been much discussed and considered as a problem for the theory of natural selection. It is therefore of great interest to understand what makes it possible. Predator psychology is an important factor in the evolution of aposematism and mimicry. In the past decades it has been suggested that certain components of prey appearance are more important to predators than others during prey assessment. We developed this idea by incorporating concepts from associative learning, and presented a new approach to explain imperfect mimicry. Our general hypothesis is that prey traits have different salience to predators. Certain traits are perceived as highly salient and are thus used primarily in the discrimination and generalization of prey, while traits with low salience are overshadowed and not used in the assessment. The salience of a trait can depend on how conspicuous or discriminable it is in the particular context, and can vary due to for example previous predator experience. We tested our ideas with wild blue tits and domestic chickens as predators, and artificial and semi-natural prey stimuli. In paper I we found that the trait that was perceived as most salient (color) was the one used to discriminate and generalize between prey. Mimics of that specific trait were highly avoided, despite differences in the other traits. We also found that salience is relative and context dependent (paper II). In a context where two traits were perceived as similarly salient, mimicry of a single trait offered intermediate protection, while mimicry of both offered high protection. In another context, the traits were perceived differently salient, and mimicry of one trait was enough for high protection. In paper III we tested a proposed scenario for the initiation of mimicry evolution in the edible butterfly mimic Papilio polyxenes asterius to its noxious model Battus philenor. The results showed that a partial similarity with the model in the salient black wing color offered intermediate protection from attacks, despite a general dissimilarity. This thesis investigates the major questions of imperfect mimicry: the initial step of mimicry evolution, the persistence of imperfect mimicry, and variation in mimic-model similarity. We conclude that mimicry evolution can begin in a non-mimetic species that acquires similarity to a model species in a high-salience trait. When multiple traits have similar salience, multi-trait mimicry is needed for higher protection. Mimicry can remain imperfect if the differences are in traits with low salience, and therefore under low or no selection pressure to change. To complete the picture, we showed that predators can have a biased generalization toward a more pronounced version of a salient trait (paper IV). The evolution of aposematism could therefore be explained by gradual enhancement of salient traits. / <p>At the time of the doctoral defense, the following papers were unpublished and had a status as follows: Paper 3: Accepted. Paper 4: Manuscript.</p>

Mule deer (Odocoileus hemionus) are an iconic species of wildlife, and populations of mule deer across much of the western U.S. have experienced recent fluctuations in size. Factors that affect the survival and subsequent recruitment of juveniles may be the preeminent cause of population fluctuations for mule deer in many areas. Many factors, including habitat loss, extreme weather, intense predation, timing and synchrony of parturition, and competition with other species may be influencing these changes. We studied two potential factors that can influence the survival of neonate mule deer in southern Utah. To better understand how predation affects mule deer, we first implemented a study of the response of mule deer to removal of coyotes in southern Utah. We monitored survival and cause-specific mortality of neonate mule deer in areas where coyotes were removed and where they were not removed. We used multi-model inference within Program MARK and a known-fate model to estimate survival of neonate mule deer in both treatments (removal and non-removal), and to investigate factors potentially influencing survival. Our results indicated that coyote control can decrease mortality and increase survival of neonate mule deer in some situations. Removal of coyotes was most effective when removal efforts occurred for multiple consecutive years, and when control efforts occurred in or near fawning habitat. Second, we examined how synchrony of parturition affects the survival and cause-specific mortality of neonate mule deer. Reproductive synchrony is a strategy that influences the survival of juveniles and the growth of populations. Our objective was to test three possible explanations for the synchrony of parturition in mule deer; 1) pressure of predation on newborns, 2) a hybrid of predation and environmental effects, and 3) weather and food availability. To determine the effects of the timing of parturition on the survival and predator-related mortality of neonate mule deer, we used multi-model inference within Program MARK and a known-fate model. Our results indicated that the timing of parturition influenced survival and predator-related mortality of neonate mule deer. There was a lag between the onset of parturition of mule deer and predation of mule deer by fawns; individuals born close to the onset of parturition had higher survival and lower predator-related mortality than those whose births were delayed relative to the onset of parturition. Since predators selected for neonate mule deer that were born late, predator learning may partially explain reproductive synchrony in mule deer. Environmental factors may have a greater effect than predation on the survival of early-born individuals.